The Foraging Tunnel System of the Namibian Desert Termite, Baucaliotermes Hainesi
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Journal of Insect Science: Vol. 10 | Article 65 Tschinkel The foraging tunnel system of the Namibian Desert termite, Baucaliotermes hainesi Walter R. Tschinkel Department of Biological Science, Florida State University, Tallahassee, FL 32303 Abstract The harvester termite, Baucaliotermes hainesi (Fuller) (Termitidae: Nasutitermitinae), is an endemic in southern Namibia, where it collects and eats dry grass. At the eastern, landward edge of the Namib Desert, the nests of these termites are sometimes visible above ground surface, and extend at least 60 cm below ground. The termites gain access to foraging areas through underground foraging tunnels that emanate from the nest. The looseness of the desert sand, combined with the hardness of the cemented sand tunnels allowed the use of a gasoline- powered blower and soft brushes to expose tunnels lying 5 to 15 cm below the surface. The tunnels form a complex system that radiates at least 10 to 15 m from the nest with cross- connections between major tunnels. At 50 to 75 cm intervals, the tunnels are connected to the surface by vertical risers that can be opened to gain foraging access to the surrounding area. Foraging termites rarely need to travel more than a meter on the ground surface. The tunnels swoop up and down forming high points at riser locations, and they have a complex architecture. In the center runs a smooth, raised walkway along which termites travel, and along the sides lie pockets that act as depots where foragers deposit grass pieces harvested from the surface. Presumably, these pieces are transported to the nest by a second group of termites. There are also several structures that seem to act as vertical highways to greater depths, possibly even to moist soil. A census of a single nest revealed about 45,000 termites, of which 71% were workers, 9% soldiers and 6% neotenic supplementary reproductives. The nest consisted of a hard outer “carapace” of cemented sand, with a central living space of smooth, sweeping arches and surfaces. A second species of termite, Promirotermes sp. nested in the outer carapace. Key Words: foraging, harvester termite, nest construction, architecture, construction Correspondence: [email protected] Associate Editor: Robert Jeanne was editor of this paper Received: 14 February 2008, Accepted: 7 July 2008 Copyright : This is an open access paper. We use the Creative Commons Attribution 3.0 license that permits unrestricted use, provided that the paper is properly attributed. ISSN: 1536-2442 | Vol. 10, Number 65 Cite this paper as: Tschinkel WR. 2010. The foraging tunnel system of the Namibian Desert termite, Baucaliotermes hainesi. Journal of Insect Science 10:65 available online: insectscience.org/10.65 Journal of Insect Science | www.insectscience.org 1 Journal of Insect Science: Vol. 10 | Article 65 Tschinkel Introduction mound and extending 25 to 30 m to the dead wood on which the termites were Termites can be roughly grouped into those feeding (Ratcliffe and Greaves 1940; Hill species that nest within their food, usually 1942; Greaves 1962). In C. lacteus, tunnels wood, and those that nest elsewhere and were more or less radial, with few cross must leave their nest in order to forage for connections, but with shafts to deeper soil. food. Of the latter type, nests may be In N. exitiosus, the radial tunnels were arboreal or subterranean, centrally located cross-connected. Hill (1925) noted or dispersed into small, connected units. subterranean passages with flattened Most termites shun the open air, and travel lumena thickly floored with “rejectamenta” to and from the foraging area by way of radiating outward from a nest of the subterranean tunnels or covered galleries. Australian Mastotermes darwiniensis, but Many species also cover the foraged he did not trace these passages far. A material with sheet galleries before dining. particularly thorough study is that of Darlington (1982), in which the Among ground-nesting termites, nests may underground foraging passages of be hidden below ground, or they may be Macrotermes michaelsoni were exposed conspicuous features of the landscape, such and quantified. as the mounds of the southern African species of Macrotermes or Trinervitermes. Many termites do not build mounds that Given a central nest, the need to forage for show above ground, but construct entirely food and an aversion toward open air, it is subterranean nests, with tunnels to the obvious that many termites must create surface. The African harvester termite, subterranean foraging tunnel systems. Such Hodotermes mossambicus, is well studied systems, however, have rarely been studied, because of occasional subterranean and are usually hardly mentioned (if at all) encounters during the digging of trenches in reviews of termite biology. Even an for construction (Hartwig 1963, 1965; authoritative treatment, such as Noirot’s Coaton and Sheasby 1975). These (1970) review of the nests of termites, gives encounters revealed that nests are located short shrift to how termites travel from their an average about 1.4 m below ground, but nests to their foraging areas. Typically, it is can be as shallow as a few cm or as deep as assumed that the termites travel in 6.7 m. Large passages connect these subterranean foraging tunnels (e.g. Sands subterranean nests to each other, and 1961), and indeed, the few existing studies smaller passages give the termites access to of subterranean foraging tunnels have the surface where they dump excavated soil revealed tunnel systems of remarkable size and forage for grass. Foraged grass is first and scale (Howse 1970; reviewed by Lee placed into small, superficial chambers for and Wood 1971). Most mound-building later transportion to the nests and species exit their nests through subterranean consumption. foraging tunnels that run a few cm below the surface. In some species, the tunnels are None of the reports on subterranean gallery short, and the termites travel some distance systems describe architectural details of the on the ground surface, but in others, the tunnels themselves or how they are tunnels may extend 25 to 30 m (or even 60 constructed. This paper reveals the intricate m) from the mound. For example, the and subtle architecture of the foraging Australian termites Coptotermes lacteus, C. tunnels of the Namibian harvester termite, brunneus, C. acinaciformis and Baucaliotermes hainesi (Fuller) Nasutitermes exitiosus constructed systems (Termitidae: Nasutitermitinae), and with 9 to 30 tunnels emanating from the describes how this complex system Journal of Insect Science | www.insectscience.org 2 Journal of Insect Science: Vol. 10 | Article 65 Tschinkel probably serves the foraging needs of the from year to year. termites. Like other harvester termites, B. hainesi foragers cut pieces of grass on the Tunnel excavation and mapping ground surface, and carry these back to Nests of B. hainesi were regularly visible at their nest. The range of this species is the surface as small mounds of cemented limited to southern Namibia and the material 10 to 15 cm high. All excavation northwestern Cape Province of South work was completed between October 22 Africa (Coaton and Sheasby 1973). and November 3, 2007. Tunnels were initially exposed by trenching around the Materials and Methods nest to locate tunnels, and excavated outward from there. The looseness of the The study site dry sand, combined with the relative The study site was located at latititude - hardness of the cemented sand tunnels 24.9702, longitude 15.9323 (according to facilitated exposure of the tunnels. The sand Google Earth) in the NamibRand Nature over the tunnels was loosened with a soft Reserve, a private reserve of about 180,000 hand broom, and the loosened sand was ha. The soil was red sand largely stabilized blown away with a gasoline-powered lawn by the grasses, Stipagrostis uniplumis blower (Husqvarna Model 356 BTx) (Video (Licht) De Winter and S. giessii Kers 1, Video 2). This process produced a (Poales: Poaceae), with circular, bare areas shallow trench 10-15 cm deep with the 5 to 15 m in diameter termed “fairy circles” mostly intact tunnels in the bottom. (van Rooyen et al. 2004), and abundant Branches and intersections were sometimes animal trails crossing it in multiple followed, but, for many branches, only the directions. The site sloped gently from initial few cm were exposed, leaving an about 1100 m elevation at the base of unknown but substantial fraction of the Jagkop mountain to about 940 m just short entire tunnel system unexposed. Tunnels of the Bushman Hills. Our two excavations that were in use were distinguished from were at approximately 1085 to 1090 m abandoned tunnels because the former elevation. This area has an arid climate remained intact upon excavation and/or where rainfall averages between 50 and 150 contained live termites mm per annum but is highly variable Figure 1. Tunnels in current use by the termites could usually be recognized by the termites found within them. Here, nasute soldiers are defending a broken tunnel. High quality figures are available online. Journal of Insect Science | www.insectscience.org 3 Journal of Insect Science: Vol. 10 | Article 65 Tschinkel when broken (Figure 1). Abandoned tunnels The dissection and collection took two tended to break, and were often filled with days. Termites from the mound and each sand. quarter of the nest from the top downwards were preserved separately. A sample of 100 In this manner, large parts of the foraging each of workers, soldiers and neotenic tunnel systems of two focal nests were supplementary reproductives (there were no exposed, one located at (lat, long) - mature alates in the nest) were killed by 24.96960, 15.93284 and the second at - freezing and air-dried for later 24.96981, 15.93403. The exposed tunnel determination of dry weight. systems were mapped by making a series of overlapping digital photographs (with a Counts were carried out in the laboratory at scale) from a uniform height (~ 1 m), like Florida State University.